Coral Reproduction on the World's Southernmost Reef at Lord Howe
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Vol. 23: 275–284, 2015 AQUATIC BIOLOGY Published online April 13 doi: 10.3354/ab00627 Aquat Biol OPENPEN ACCESSCCESS Coral reproduction on the world’s southernmost reef at Lord Howe Island, Australia Andrew H. Baird1,*, Vivian R. Cumbo1, Sallyann Gudge2, Sally A. Keith1,3, Jeffrey A. Maynard4,5, Chun-Hong Tan1,6, Erika S. Woolsey1 1ARC Centre of Excellence for Coral Reef Studies, James Cook University, Townsville, QLD 4811, Australia 2NSW Department of Primary Industries, Lord Howe Island Marine Park, PO Box 161, Lord Howe Island, NSW 2898, Australia 3Center for Macroecology and Evolution, University of Copenhagen, Universitetsparken 15, 2100 Copenhagen, Denmark 4Department of Ecology and Evolutionary Biology, Cornell University, Ithaca, NY 14853, USA 5Laboratoire d’Excellence «CORAIL» USR 3278 CNRS−EPHE, CRIOBE, Papetoai, Moorea, French Polynesia 6School of Marine Science and Environment, Universiti Malaysia Terengganu, 21030 Kuala Terengganu, Terengganu, Malaysia ABSTRACT: Despite a recent expansion in the geographic extent of coral reproductive research, there remain many regions in the Indo-Pacific where knowledge is limited. For example, Lord Howe Island is the southernmost reef system in the world (31° S); however, very little is known of the repro- ductive biology of the coral fauna. Here, aspects of the reproductive biology and the timing of repro- duction for 40 of the approximately 65 species that occur on Lord Howe Island are documented. In December 2010, field assessments of the stage of gamete maturity in Acropora spp. colonies sug- gested that 5 species spawned in December 2010 and 11 in January 2011. In January 2012, similar sampling suggested that 12 Acropora species spawned in January and 1 in February. In addition, 11 species from 10 genera broadcast spawned gametes from 17:30 to 24:00 h in January 2012, 10 to 12 d after full moon. Goniastrea favulus was inferred to spawn prior to 17:00 h, 6 to 12 d after full moon and Porites heronensis released brooded larvae. The reproductive biology of 3 other brooding species was examined using dissections and histology monthly for 1 yr from April 2011. Of these, Se- riatopora hystrix contained planulae between November 2011 and March 2012, Stylophora pistillata contained planulae between November 2011 and February 2012. No eggs or planulae were ob - served in Pocillopora damicornis. In conclusion, the spawning patterns on Lord Howe Island are consistent with other locations in the Indo-Pacific: multi-species synchronous spawning episodes occur after full moons, when water temperatures are relatively high. KEY WORDS: Coral reefs · Larval ecology · Spawning timing · Egg size INTRODUCTION often context-dependent suggests we should seek to quantify reproductive phenology across a wide geo- Coral reproductive research in the Indo-Pacific has graphic extent. Whilst the geographic focus of coral been concentrated historically in regions close to reproductive research has recently expanded to in - marine research stations in developed countries (see clude more remote regions such as India (Raj & review by Baird et al. 2009b). However, coral repro- Edward 2010), New Caledonia (Baird et al. 2010), the duction is increasingly recognised as a fundamental Persian Gulf (Bauman et al. 2011) and Yemen (Baird process to maintain ecosystem function within coral et al. 2014), reproductive phenology on marginal reefs and, as such, interest in improving our un- reefs at the extreme geographical limits of coral dis- derstanding of this process has grown rapidly. In tributions has yet to be quantified. Corals on mar- addition, recognition that ecological processes are ginal reefs would be expected to be the least con - © The authors 2015. Open Access under Creative Commons by *Corresponding author: [email protected] Attribution Licence. Use, distribution and reproduction are un - restricted. Authors and original publication must be credited. Publisher: Inter-Research · www.int-res.com 276 Aquat Biol 23: 275–284, 2015 sistent with a broader pattern and, therefore, offer 2009) and therefore replenishment is likely to be potential to determine the limits of reproductive highly dependent on local reproductive output or phenology. irregular input of propagules from distant upstream The increased geographical extent across which re- reefs (Fellegara et al. 2013, Madsen et al. 2014). productive phenology has been quantified has altered Most broadcast-spawning colonies spawn only once our understanding of coral reproductive synchrony per year (Harrison & Wallace 1990, Baird et al. 2009b). and the processes that regulate the timing of coral In contrast, oogenic cycles are much shorter in corals reproduction (Baird et al. 2009b, van Woesik 2010). that brood larvae (e.g. Stoddart & Black 1985, Per- For example, highly synchronous spawning, both mata et al. 2000), and consequently individual colonies within and among species, is not restricted to higher can release larvae multiple times per year. For exam- latitude reefs as earlier hypothesized (Oliver et al. ple, individual colonies of Pocillopora damicornis, 1988). In particular, multi-species synchronous spawn- Seriatopora hystrix and Stylophora pistillata release ing events (sensu Willis et al. 1985) are now de - larvae throughout the year at Palau, whereas at scribed for >25 locations globally (Baird et al. 2009b, Heron Island, on the southern Great Barrier Reef, Kongjandtre et al. 2010, Bouwmeester et al. 2011, breeding is seasonal (see review in Tanner 1996). Permata et al. 2012). In addition, coral reproduction Similarly, Isopora palifera releases planulae through- in most regions is highly seasonal, with the vast out the year in Papua New Guinea, but only in the majority of reproductive activity concentrated in 2 to summer months on Heron Island (Kojis 1986). 3 months each year (Baird et al. 2009a). This trend The highly seasonal nature of coral reproduction holds in equatorial locations (e.g. Singapore: Guest has important implications for reef ecology and con- et al. 2005; e.g. Kenya: Mangubhai & Harrison 2008) servation (Guest 2008). Numerous reef organisms, and in regions where breeding patterns had previ- including many fishes, time their reproductive cycles ously been described as aseasonal (e.g. the Red Sea: to benefit from this seasonal abundance of nutrients Shlesinger & Loya 1985) due to incomplete sampling (Pratchett et al. 2001, McCormick 2003). In addition, (Hanafy et al. 2010, Bouwmeester et al. 2015). knowledge of coral reef phenology allows the use of Despite this recent expansion in the geographic temporal management strategies that can mitigate extent of studies on coral reproduction, there remain the effects of coastal development on coral reefs many regions in the Indo-Pacific where research is (Richmond 1997). For example, dredging or discharge limited. For example, Lord Howe Island contains the of liquid waste from heavy industry can be prohibited southernmost reef system in the world (31.53° S, during coral spawning, ensuring that the seasonal 159.08° E); and is a World Heritage-listed marine pro- production of propagules for reef replenishment is tected area in the Tasman Sea, with highly distinc- not jeopardized (Baird et al. 2011). tive marine fauna including 9 endemic fish species In this study, we document the day or month of (Francis 1993) and 47 endemic species of algae (MPA propagule release for 34 of the approximately 65 coral 2010). Lord Howe Island supports a scleractinian species on Lord Howe Island. We also document fauna of approximately 65 species (Veron 1993); aspects of the reproductive biology of many of these however, almost nothing is known of the reproduc- species, including the sexual system, mode of larval tive biology of these corals, in particular the timing of development, transmission of Symbiodinium spp. and spawning and the length of the reproductive season. egg size on release. Goniastrea favulus was observed broadcast spawn- ing at approximately 16:30 h ‘during mid-January, 1977’ (Kojis & Quinn 1981), 6 Acropora spp. and MATERIALS AND METHODS Cyphastrea microphthalma were observed setting (immediately prior to spawning) or spawning 8 to 9 Reproductive condition of Acropora species nights after the full moon, and Isopora cuneata plan- ulated 9 nights after the full moon in January 2007 Acropora were sampled 8 to 10 d before full moon (Harrison 2008). In addition, local tourist operators on 21 December 2010, and 3 to 5 d after full moon on often run nighttime snorkelling trips to observe 9 January 2012. These sampling times were chosen corals spawning following the full moon in February because previous work on Lord Howe Island indi- (P. Busteed pers. comm.). The reproductive bio logy cated that Acropora spp. spawn 8 to 10 d after full of high-latitude corals, such as those on Lord Howe moon (Harrison 2008). Twenty-three Acropora spp. Island, is of particular interest, because these popula- were sampled to determine their reproductive condi- tions are often geographically isolated (Noreen et al. tion (for full list of species see Table 1). Three repro- Baird et al.: Coral reproduction on Lord Howe Island 277 under the jetty during daylight hours (for species list see Table 2). Colonies were submerged in buckets and transported to the research station each day at 17:30 h, where they were maintained until 08:00 h the following morning, with 1 complete seawater change at 24:00 h. Colonies were inspected approxi- mately ev ery 30 min while at the research sta tion to capture the onset